Outline the events of sequence which lead to the formation

1. A mature xylem vessel Xylem, in botany, part of the vascular system that conveys water and dissolved minerals from the roots to the rest of the plant and may also furnish mechanical support. Xylem consists of specialized waterconducting tissues made up mostly of narrow, elongated, hollow cells. These cells may be of several types, including tracheid (the basic cell type), vessel members, fibers, and parenchyma. Xylem constitutes the major part of a mature woody stem or root; the wood of a tree is composed of xylem. Xylem formation begins when the actively dividing cells of growing root and shoot tips (apical meristems) give rise to primary xylem. As the growing part of the plant builds past the xylem thus formed, the vascular cambium produces secondary xylem tissues that cover the primary xylem. When this happens the primary xylem cells become dead and empty, losing their conducting function and forming a hard skeleton that serves only to support the plant. Thus, in the trunk and older branches of a large tree only the outer part of the wood (secondary xylem) serves in water conduction, while the inner part (heartwood) is composed of dead but structurally strong primary xylem. .

2. A mature sieve tube Unlike the elements of the xylem, which undergo a complete and total breakdown during differentiation and transport occurs through dead, empty cells; sieve elements are living cells, with cytoplasm, at functional maturity. The term sieve refers to the clusters of pores, the sieve areas, through which the cytoplasm of adjacent elements are interconnected. Most of the sieve areas are concentrated on the overlapping ends of the long sieve tube elements. The cytoplasm of sieve elements is unique, however. As the sieve element differentiates and matures, it undergoes a selective breakdown. The two major changes are the

breakdown of the nucleus and the vacuolar membrane, but other changes occur as well: ribosomes, Golgi apparatus, and the cytoskeleton are lost, and all the remaining components become distributed along the wall. The plasma membrane, smooth endoplasmic reticulum, some plastids, and mitochondria remain. Some sieve-tube elements possess a proteinaceous substance called P-protein, which also becomes distributed along the wall. In cut sections of phloem tissues, slime plugs of P-protein are usually found near the sieve plates. Slime plugs are believed to result from the surging of the contents of sieve tubes that are severed. Some botanists believe that P-protein helps seal the sieve-plate pores at the time of wounding.

Differentiation of a sieve-tube element: (a) The mother cell of the sieve-tube element undergoing division. (b) Division has resulted in formation of a young sieve-tube element and a companion cell. After division, one or more P-protein bodies arise in the cytoplasm, which is separated from the vacuole by a tonoplast. The wall of the young sieve-tube element has thickened, and the sites of the future sieveplate pores are represented by plasmodesmata. Each plasmodesma is now surrounded by a platelet of callose on either side of the wall. (c) The nucleus is degenerating, the tonoplast is breaking down, and the P-protein bodies are dispersing in the cytoplasm lining the wall. At the same time, the plasmodesmata of the developing sieve plates are beginning to widen into pores. (d) At maturity, the sieve-tube element lacks a nucleus and a vacuole. All of the remaining protoplasmic components, including the P-protein, line the walls, and the sieve-plate pores are open. The callose platelets were removed as the pores widened. Not shown here but also present in the mature sievetube element are smooth endoplasmic reticulum, mitochondria, and plastids.

What are the structural and physiological differences between xylem vessel and phloem?

By structure, phloem has an elongated, tubular shape with thin walled sieve tubes. The sieve tubes have pores at each end in the cross walls and microtubules that extend between sieve elements allowing longitudinal flow of material whereas, xylem has a tubular shape with no cross walls which allows a continuous column of water and facilitated more transport within the xylem vessel. There are two types, protoxylem (first formed) and metaxylem (mature xylem). Phloem sieve tube is made up of living cells that function all the cell metabolic reactions, while the xylem vessel is made up of living cells which probably dont work. The phloems cell wall material is made of cellulose with a lining cytoplasm which contains all cell organelles, while the xylem cell wall material is lignified and cemented that stops it from being broken by high water potential. In addition to that, xylem vessel is impermeable due to the fact that is cemented, while phloem is permeable meaning it allows the entrance of materials and allows them to leave as well. Physiologically, the elements within the xylem and phloem can act independently of one another, yet phloem relies on water provided by xylem in order to provide driving forces required for long distances translocation of assimilated material.

By Sylvia Kalisa